A systematic revision of Baconia Lewis (Coleoptera, Histeridae, Exosternini) Author Caterino, Michael S. Author Tishechkin, Alexey K. text ZooKeys 2013 343 1 297 http://dx.doi.org/10.3897/zookeys.343.5744 journal article http://dx.doi.org/10.3897/zookeys.343.5744 1313-2970-343-1 Baconia Lewis, 1885: 462 Type species: Baconia loricata Lewis, 1885: 463, designated by Bickhardt 1917 : 163. Binhister Cooman, 1934: 122 (type species Binhister barbarus Cooman, by original designation); previously recognized as a subgenus, it is here fully synonymized; comb. n. Description. Size range: Length 1.0-5.0mm; width 0.6-4.0mm; Body: ovoid to elongate, sides broadly rounded to sub- or fully parallel, convex to very strongly flattened; color rufescent to frequently piceous or metallic; glabrous or rarely finely setose. Head: frons convex, flat, or deeply depressed, frontal stria usually present along inner margin of eyes, variably interrupted or obsolete across front, frons and epistoma frequently separated by weak to strong transverse carina; supraorbital stria present or absent; epistoma depressed to flat or convex, frequently swollen along apical margin, apical margin usually straight; labrum usually much wider than long, up to 4 x or more, usually emarginate apically, but may be straight, bisinuate, or weakly produced; antennal scape usually short, stout, only weakly expanded to apex (Fig. 1A), may be longer, and/or expanded apically; antennal club generally completely tomentose, though rarely glabrous basally, annuli absent, but with 4 characteristic sensory slits on upper and lower surfaces (Fig. 1B), rarely with additional subapical sensorial patch (Fig. 54E); submentum angulate at base, truncate to projecting along distal margin, with few simple setae; gular sutures finely impressed, extending anterolaterad, uninterrupted to basal corner of buccal cavity; mentum subquadrate, sides weakly convergent, apical margin truncate to weakly emarginate, bearing few simple setae; labium with palpifers prominent, palpi with three palpomeres, the basalmost very short, the distal two with short, scattered setae; maxilla with cardo short, transverse, glabrous, stipes triangular, bearing few simple setae, palpi with four palpomeres, the basalmost very short; mandibles (Figs 1A, 3A, 28C) generally each with basal tooth, may be blunt or strong and acute, mandible frequently furrowed along lower, outer edge, may have ventral (mesal) pore and associated (presumed) secretory channel (Fig. 1C). Pronotum: sides parallel to convergent apically; marginal stria usually present and continuous around lateral and anterior margins; lateral submarginal stria present or absent; anterior corners nearly always weakly depressed (Fig. 1D); prescutellar impression absent; disk with single pair of anterior marginal gland openings, usually located close to anterior margin, behind eye on each side, discal punctation highly varied. Elytra: with 2-3 epipleural striae, outer subhumeral stria rarely present, inner subhumeral stria frequently present, often restricted to short basal fragment, dorsal striae 1-5 and sutural stria highly varied, variously abbreviated from base or apex or entirely absent; elytral disk nearly always with distinct secondary punctures in apical half or less (Fig. 1E). Prosternum: prosternal keel varied in width, often quite broad in depressed species, basal margin varied from emarginate to truncate, rarely outwardly arcuate; carinal striae generally present, usually complete, free, rarely abbreviated anteriorly, united or obsolete ; prosternal lobe short to moderate in length, apical margin subtruncate to broadly arcuate, rarely bisinuate; marginal stria of prosternal lobe usually distinct across middle, variably obsolete at sides. Mesoventrite: anterior mesoventral margin ranging from distinctly emarginate to distinctly projecting, marginal stria complete to absent, rarely with secondary submarginal stria; mesometaventral stria usually present, most frequently arched forward onto mesoventrite, may in some cases partially displace or completely replace marginal mesoventral stria. Metaventrite: Anterior margin, i.e., mesometaventral suture, frequently arched forward (mirrored in most, but not all cases, in mesometaventral stria), inner lateral metaventral stria generally present, extending from near inner corner of mesocoxa toward metacoxa, or toward posterior corner of metepisternum in the most depressed species, variably sinuate or ab breviated apically; outer lateral metaventral stria present or absent; metaventral disk usually coarsely punctate at sides, impunctate at middle. Abdomen: 1st abdominal ventrite with one or two lateral striae along inner edge of metacoxa, disk usually impunctate at middle, but with conspicuous median punctures in various species; abdominal ventrites 2-5 usually punctate at sides, rarely with dense punctures extending across middle of disk (Fig. 43F). Propygidium often with basal transverse stria (Fig. 5E), disk variably punctate sexually dimorphic in one species; single pair of propygidial gland openings usually conspicuous (Fig. 1F), situated on each side variably near basal margin; pygidium never with apical marginal stria, usually densely punctate, very rarely sexually dimorphic, with male's setose or otherwise modified; both sexes bearing pygidial trichomes in one species.Legs: Protibia usually rather narrow, with 0-5 unevenly spaced marginal teeth (Fig. 2A), the outer margin nearly always finely serrulate along entire length (Fig. 2A); protibial spurs present, usually short, weakly curved; mesotibia usually with 1-2 weak marginal spines (Fig. 2B), rarely lacking spines; metatibia rarely with any marginal spines (Fig. 2C), generally smooth, with outer apical corner slightly prolonged; tarsi not obviously dimorphic, tarsomeres 1-4 short, usually bearing only single pair of apical setae, tarsomere 5 about as long as 2-4 together, usually weakly dorsoventrally curved (Fig. 2D); tarsal claws simple, separate. Male genitalia:Accessory sclerites absent. T8 generally short, broad, with basal rim strongly sclerotized, basolateral edge extending beneath to inner corner of ventrolateral apodeme; ventrolateral apodeme usually acute, with distal portion strongly reduced, T8 usually broadly open beneath; basal emargination very shallow to moderately deep; basal membrane attachment line rarely evident, usually intersecting basal emargination; distal margin weakly sclerotized, poorly defined, usually vaguely emarginate. S8 articulated at basal corners with ventrolateral apodemes of T8, apical guides weakly developed, halves separate or fused; if separate, halves usually strongly divergent from base, with apices narrow, often rounded, bare to conspicuously setose; if halves of S8 fused, apical margin usually weakly emarginate, frequently with apicoventral velar membrane, bare to conspicuously setose. T9 usually divided, rarely united, with basal apodemes long and slender to short and broad; ventrolateral apodemes weak to strong, opposing or recurved basad, very rarely fused beneath; distal apices usually weakly opposed, subacute to truncate; subapical seta often present on sides. T10 entire, weakly sclerotized, apical margin rounded to weakly emarginate. S9 usually desclerotized along midline, rarely entirely divided; stem very narrow to moderately broad, frequently with ventral keel along much of length; stem rarely absent, with entire T9 short, subcordate; head of S9 usually broad, acute apicolaterally, with apical margin shallowly emarginate to sinuate; tegmen relatively simple, shallowly incised apically, parallel-sided to tapered apically, lacking medioventral tooth or process; median lobe narrow, simple, in a few species associated with small, articulated apical denticulate plates (Fig. 41O); basal piece usually short, with superficial membrane attachment line and oval, asymmetrical basal foramen. Female genitalia: T8 forming a single plate, apical margin usually emarginate; S8 entire or with median plate isolated, with basal baculi detached, articulated with sternites, basally subparallel; S9 usually present, elongate; median coxite articula tion present; valviferae paddle-shaped; coxites varied in shape, subquadrate to elongate, with 2-5 apical marginal teeth, with distinct, articulated apical stylus; bursa copulatrix usually completely membraneous, rarely with small sclerotizations of bursal wall; generally with single, bulbous, weakly sclerotized spermatheca, inserted near or at apex of bursa copulatrix; single, basically thin and elongate spermathecal gland present, generally attached near midpoint of spermatheca. Figure 1. Baconia generic characters. A Frons of Baconia anthracina B Antenna of Baconia tricolor C Mouthparts of Baconia gibbifer D Pronotum of Baconia disciformis E Elytron of Baconia anthracina F Pygidia of Baconia tricolor . Figure 2. Baconia generic characters. A Protibia of Baconia tricolor B Mesotibia of Baconia tricolor C Metatibia of Baconia anthracina D Mesotarsus of Baconia insolita . Diagnosis. Although initially characterized on the basis of extreme flattening of the body, the species of Baconia in fact span a broad range from convex to very flat. They are frequently metallic in appearance, although many species exibit no hint of metallic coloration. They are best recognized by their unique antennal club sensoria - in all species the annuli are reduced to a set of 4 distinct sensory slits on both upper and lower surfaces (Figs 1B, 54E, 64C). These are found in no other Histeridae . Additional characters that can help to diagnose the genus if the antennal club is not visible: antennal scape usually short (Fig. 1A), no more than twice as long as wide, its apex obliquely truncate; frontal stria rarely complete; inner margins of eyes often strongly convergent dorsally (Fig. 1A); both mandibles usually with distinct basal tooth (Fig. 1A); anterior corners of pronotum concave to depressed (Fig. 1D); antescutellar fovea absent; lateral pronotal stria, when present, close to margin and often carinate; elytral disk with secondary punctures near apex (or, if generally punctate, punctures becoming larger and denser toward apex; Fig. 58A); prosternal striae usu ally present, rarely meeting anteriorly; protibia usually finely serrulate between major marginal spines; metatibia (and often mesotibia) with few or no marginal spines, and frequently prolonged at outer apex (Fig. 2C); and apical tarsomere long and somewhat curved dorsoventrally (Fig. 2D). In the Neotropical region, the depressed anterior pronotal corners, minimially spinose posterior tibiae, and presence of apical elytral secondary punctures will distinguish them from all other histerids in the region. Hypobletus spp. may show comparable flatness to some Baconia , but are always rufescent in color and parallel-sided in body form. The most flattened Baconia spp. are nearly all metallic and/or rounded at the sides. In the Oriental region, where all the Baconia spp. are moderately to strongly depressed in body shape, there may be some Platysomatini that are generally similar in body form, but these will always exhibit S-shaped protarsal grooves and complete antennal annuli. Checklist of the species of Baconia We establish here a series of informal species groups. These generally correspond to groups that we feel reflect monophyletic groups within the genus (and are supported as such in preliminary analyses). However, as much as anything they are intended to facilitate identification and description, uniting species that are similar and appear related. Ordering of species within species groups is also intended to reflect phylogeny to a certain degree, facilitating comparisons among closely related species and their diagnoses. Baconia loricata group Baconia loricata Lewis, 1885 Baconia patula Lewis, 1885 Baconia gounellei (Marseul, 1887a) Baconia jubaris (Lewis, 1901) Baconia festiva (Lewis, 1891) Baconia foliosoma sp. n. Baconia sapphirina sp. n. Baconia furtiva sp. n. Baconia pernix sp. n. Baconia applanatis sp. n. Baconia disciformis sp. n. Baconia nebulosa sp. n. Baconia brunnea sp. n. Baconia godmani group Baconia godmani (Lewis, 1888) Baconia venusta (J. E. LeConte, 1845) Baconia riehli (Marseul, 1862), comb. n. Baconia scintillans sp. n. Baconia isthmia sp. n. Baconia rossi sp. n. Baconia navarretei sp. n. Baconia maculata sp. n. Baconia deliberata sp. n. Baconia excelsa sp. n. Baconia violacea (Marseul, 1853) Baconia varicolor (Marseul, 1887b) Baconia dives (Marseul, 1862) Baconia eximia (Lewis, 1888) Baconia splendida sp. n. Baconia jacinta sp. n. Baconia prasina sp. n. Baconia opulenta sp. n. Baconia illustris (Lewis, 1900) Baconia choaspites (Lewis, 1901) Baconia lewisi Mazur, 1984 Baconia salobrus group Baconia salobrus (Marseul, 1887b) Baconia turgifrons sp. n. Baconia crassa sp. n. Baconia anthracina sp. n. Baconia emarginata sp. n. Baconia obsoleta sp. n. Baconia ruficauda group Baconia ruficauda sp. n. Baconia repens sp. n. Baconia angusta group Baconia angusta Schmidt, 1893a Baconia incognita sp. n. Baconia guartela sp. n. Baconia bullifrons sp. n. Baconia cavei sp. n. Baconia subtilis sp. n. Baconia dentipes sp. n. Baconia rubripennis sp. n. Baconia lunatifrons sp. n. Baconia aeneomicans group Baconia aeneomicans (Horn, 1873) Baconia pulchella sp. n. Baconia quercea sp. n. Baconia stephani sp. n. Baconia irinae sp. n. Baconia fornix sp. n. Baconia slipinskii Mazur, 1981 Baconia submetallica sp. n. Baconia diminua sp. n. Baconia rufescens sp. n. Baconia punctiventer sp. n. Baconia aulaea sp. n. Baconia mustax sp. n. Baconia plebeia sp. n. Baconia castanea sp. n. Baconia lescheni sp. n. Baconia oblonga sp. n. Baconia animata sp. n. Baconia teredina sp. n. Baconia chujoi (Cooman, 1941) Baconia barbarus (Cooman, 1934) Baconia reposita sp. n. Baconia kubani sp. n. Baconia wallacea sp. n. Baconia bigemina sp. n. Baconia adebratti sp. n. Baconia silvestris sp. n. Baconia cylindrica group Baconia cylindrica sp. n. Baconia chatzimanolisi sp. n. Baconia gibbifer group Baconia gibbifer sp. n. Baconia piluliformis sp. n. Baconia maquipucunae sp. n. Baconia tenuipes sp. n. Baconia tuberculifer sp. n. Baconia globosa sp. n. Baconia insolita group Baconia insolita (Schmidt, 1893a), comb. n. Baconia burmeisteri (Marseul, 1870) Baconia tricolor sp. n. Baconia pilicauda sp. n. Baconia riouka group Baconia riouka (Marseul, 1861) Baconia azuripennis sp. n. Baconia famelica group Baconia famelica sp. n. Baconia grossii sp. n. Baconia redemptor sp. n. Baconia fortis sp. n. Baconia longipes sp. n. Baconia katieae sp. n. Baconia cavifrons (Lewis, 1893), comb. n. Baconia haeterioides sp. n. Baconia micans group Baconia micans (Schmidt, 1889a) Baconia carinifrons sp. n. Baconia fulgida (Schmidt, 1889c) Baconia incertae sedis Baconia chilense (Redtenbacher, 1867) Baconia glauca (Marseul, 1884) Baconia coerulea (Bickhardt, 1917) Baconia angulifrons sp. n. Baconia sanguinea sp. n. Baconia viridimicans (Schmidt, 1893b) Baconia nayarita sp. n. Baconia viridis sp. n. Baconia purpurata sp. n. Baconia aenea sp. n. Baconia clemens sp. n. Baconia leivasi sp. n. Baconia atricolor sp. n. Key to species of Baconia

Figs 49H50BFigs 49E50A
Baconia chujoi
Fig . 49E	Baconia reposita
Fig. 73C	Baconia glauca
Fig. 49H
st	Baconia kubani
Fig. 50Est	Baconia adebratti
Fig. 50G
Baconia barbarus
Baconia silvestris
Baconia wallacea
Baconia bigemina
Fig. 69A	Baconia cavifrons
Fig. 69F	Baconia haeterioides
th	Baconia loricata
th
Fig. 5E
Fig. 3C	Baconia patula
thFig. 3E	Baconia gounellei
th	Baconia nebulosa
Baconia loricata
Baconia applanatis
Baconia brunnea
Fig. 9A-B	Baconia disciformis
Fig. 5A	Baconia jubaris
Fig. 54F	Baconia gibbifer
Fig. 54Cst	Baconia gibbifer
Fig. 54G
Fig. 54H
thFig. 56A	Baconia tenuipes
Fig. 54G	Baconia piluliformis
Fig. 54H	Baconia maquipucunae
nd
thFig. 56CFig. 56B	Baconia tuberculifer
thFig. 56D	Baconia globosa
Figs 3052th	Baconia cylindrica ruficauda
thth
Fig. 30Ath	Baconia ruficauda
th
th	Baconia teredina
th
th	Baconia chatzimanolisi
th	Baconia cylindrica
Figs 32A-D, E, G35Cth	Baconia angusta
th
Fig. 32H	Baconia bullifrons
Fig. 35C	Baconia rubripennis
Fig. 35B	Baconia dentipes
Figs 33E, Kth
th
Fig. 33EFig. 33D	Baconia angusta
Fig. 33KFig. 33J	Baconia incognita
Figs 36E-F
Fig . 33N
thFig. 37CFigs 37E-F	Baconia lunatifrons
thFig. 36CFigs 36I-J	Baconia subtilis
Figs 33O-P	Baconia guartela
Figs 34M-N	Baconia cavei
Figs 5860	Baconia insolita
Fig. 58C	Baconia burmeisteri
Fig. 58A	Baconia insolita
Fig. 61E	Baconia pilicauda
Fig. 60BFig. 61D	Baconia tricolor
Fig. 68AFig. 68B	Baconia katieae
Fig. 16A	Baconia maculata
Figs 64A, E, G	Baconia famelica
Fig . 66CFig. 66A	Baconia longipes
Fig. 64E	Baconia redemptor
Fig. 64B	Baconia famelica
Baconia grossii
Fig. 62F	Baconia azuripennis
Fig. 73D	Baconia coerulea
Baconia violacea
Fig. 71D	Baconia carinifrons
Baconia micans
Fig . 38C	Baconia pulchella
Fig. 38A	Baconia aeneomicans
Baconia quercea
Fig. 40FFig. 41F	Baconia submetallica
Fig. 40BFig. 40AFig. 39P	Baconia irinae
Fig. 16C	Baconia deliberata
Fig. 11A	Baconia godmani
Fig. 16E	Baconia excelsa
Fig. 82CFig. 82D	Baconia aenea
Fig. 71EFig. 71Fth	Baconia fulgida
Figs 11 G-H	Baconia riehli
ththFig. 6D	Baconia pernix
Fig. 6C	Baconia furtiva
Fig. 5G	Baconia foliosoma
Fig. 11F	Baconia isthmia
Figs 6 A-B	Baconia sapphirina
Figs 5 C-D	Baconia festiva
Fig. 22D	Baconia lewisi
Fig. 11D	Baconia venusta
Fig. 18G	Baconia jacinta
Fig. 80C	Baconia viridis
th
Fig. 21E	Baconia illustris
Baconia navarretei
Fig. 80B	Baconia nayarita
Fig. 18D	Baconia eximia
thth
Fig. 30B	Baconia repens
Fig. 18EFig. 18F	Baconia splendida
Fig. 14C	Baconia rossi
Fig. 22A	Baconia choaspites
Baconia scintillans
Fig. 82EFig. 82F	Baconia clemens
thFig. 21A	Baconia prasina
th
Fig. 77B	Baconia angulifrons
Fig . 16F	Baconia violacea
Fig. 18BFig. 19D	Baconia dives
Fig. 18AFig. 19B	Baconia varicolor
Figs 77F-G	Baconia viridimicans
Fig. 82A	Baconia purpurata
Fig. 21D	Baconia opulenta
Fig. 64HFig. 64G	Baconia fortis
thFig. 73A	Baconia chilense
Fig. 62A	Baconia riouka
Fig. 38F	Baconia stephani
thFig. 84A	Baconia leivasi
Figs 40A, Cth
th
Fig . 47C
Fig. 77C	Baconia sanguinea
Fig. 47C	Baconia lescheni
Fig. 47F	Baconia animata
Fig. 47D	Baconia oblonga
Baconia rufescens
Figs 40B43F
Fig. 43F	Baconia castanea
Fig. 40H	Baconia diminua
Fig. 40D	Baconia fornix
Fig. 43E	Baconia mustax
Baconia submetallica
Baconia plebeia
Baconia slipinskii
Fig. 42M	Baconia aulaea
Fig. 42P	Baconia punctiventer
thFig. 24A
th
Fig. 84C	Baconia atricolor
th
Fig. 24C	Baconia salobrus
Fig. 28C	Baconia emarginata
Fig. 28A	Baconia anthracina
Baconia obsoleta
Fig. 26A
Fig. 26A	Baconia turgifrons
Fig. 26C	Baconia crassa